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E Covey

Publications and source records attributed to E Covey.

33 records · Page 2Linked to original sources
PubMed1994

Neural tuning for sound duration: role of inhibitory mechanisms in the inferior colliculus.

Duration is a biologically important feature of sound. Some neurons in the inferior colliculus of the big brown bat, Eptesicus fuscus, are tuned to sound duration, but it is unclear at what level the tuning originates or what neural mechanisms are responsible for it. The application of antagonists of the inhibitory neurotransmitters gamma-aminobutyric acid or glycine to neurons in the inferior colliculus eliminated duration tuning. Whole-cell patch-clamp recordings of synaptic currents suggested that inhibition produces a temporal frame within which excitation can occur. A model is proposed in which duration tuning arises when an early, sustained inhibitory input interacts with a delayed, transient excitatory input.

Acoustic Stimulation↗
PubMed1994

Frequency tuning and response latencies at three levels in the brainstem of the echolocating bat, Eptesicus fuscus.

To determine the level at which certain response characteristics originate, we compared monaural auditory responses of neurons in ventral cochlear nucleus, nuclei of lateral lemniscus and inferior colliculus. Characteristics examined were sharpness of frequency tuning, latency variability for individual neurons and range of latencies across neurons. Exceptionally broad tuning curves were found in the nuclei of the lateral lemniscus, while exceptionally narrow tuning curves were found in the inferior colliculus. Neither specialized tuning characteristic was found in the ventral cochlear nuclei. All neurons in the columnar division of the ventral nucleus of the lateral lemniscus maintained low variability of latency over a broad range of stimulus conditions. Some neurons in the cochlear nucleus (12%) and some in the inferior colliculus (15%) had low variability in latency but only at best frequency. Range of latencies across neurons was small in the ventral cochlear nucleus (1.3-5.7 ms), intermediate in the nuclei of the lateral lemniscus (1.7-19.8 ms) and greatest in the inferior colliculus (2.9-42.0 ms). We conclude that, in the nuclei of the lateral lemniscus and in the inferior colliculus, unique tuning and timing properties are built up from ascending inputs.

Animals↗
PubMed1993

Response properties of single units in the dorsal nucleus of the lateral lemniscus and paralemniscal zone of an echolocating bat.

1. Connectional evidence suggests that the dorsal nucleus of the lateral lemniscus (DNLL) and the paralemniscal zone (PL) function as centers for binaural analysis interposed between the superior olivary complex and the midbrain. In addition, the DNLL is known to be a major source of inhibitory input to the midbrain. The aim of this study was to characterize the response properties of neurons in DNLL and PL of the echolocating bat Eptesicus fuscus, a species that utilizes high-frequency hearing and that might be expected to have a large proportion of neurons responsive to interaural differences in sound level. 2. Auditory stimuli were presented monaurally or binaurally to awake animals, and responses of single units were recorded extra-cellularly with the use of glass micropipettes. 3. Below the ventrolateral border of the inferior colliculus is a region that contains large gamma-aminobutyric acid-positive neurons. On the basis of its immunohistochemical reactivity, this entire region could be considered as DNLL. However, within the area, there was an uneven distribution of binaural responses. Caudally, binaural neurons made up 84% (41/49) of those tested, but rostrally only 29% (6/21). For this reason the rostral area is considered as a separate functional subdivision and referred to as the dorsal paralemniscal zone (DPL). PL is located ventral to DPL and medial to the intermediate and ventral nuclei of the lateral lemniscus; in PL 88% (14/16) of neurons were binaural. 4. Most neurons responded only to a contralateral stimulus when sounds were presented monaurally. Out of 49 neurons in DNLL, 42 responded only to a contralateral sound, 1 responded only to an ipsilateral sound, and 6 responded to sound at either ear. In the DPL, all of the 21 neurons tested responded to a contralateral sound and none to an ipsilateral sound. Out of 16 neurons in the PL, 11 responded only to a contralateral sound, 1 responded only to an ipsilateral sound, and 4 responded to sound at either ear. 5. When sounds were presented at both ears simultaneously, several different patterns of binaural interaction occurred. The most common pattern was suppression of the response to sound at one ear by sound at the other ear. In DNLL, 57% (28/49) of neurons showed this type of binaural interaction. Another 10% (5/49) showed facilitation at some interaural level differences and suppression at others, and another 10% (5/49) showed facilitation at some interaural level differences but no suppression.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1992

Monaural interaction of excitation and inhibition in the medial superior olive of the mustached bat: an adaptation for biosonar.

In most mammals, the superior olive is the first stage for binaural interaction. Neurons in the medial superior olive (MSO) receive excitatory input from both ears and are sensitive to interaural time or phase differences of low-frequency sounds. The mustached bat (Pteronotus parnellii parnellii), a small echolocating species with high-frequency hearing, probably does not use interaural time or phase differences as cues for sound localization. Although the mustached bat has a large MSO, there is some evidence that it is functionally different from the MSO in nonecholocating mammals. Most MSO neurons in the mustached bat are monaural, excited by a contralateral sound. Their responses are phasic and correlated with either the onset or the offset of a sound. As a first step in determining the origin of these phasic monaural responses, we traced the connections of the MSO by using both retrograde and anterograde transport methods. Excitatory inputs to the MSO originate from spherical cells in the anteroventral cochlear nucleus, almost exclusively from the contralateral side. Glycinergic inhibitory input is relayed from the contralateral cochlear nucleus through the medial nucleus of the trapezoid body. To investigate the interactions of the contralateral excitatory and inhibitory inputs at the level of the MSO cell, we recorded sound-evoked responses and applied glycine or its antagonist by using microiontophoresis. The results show that the phasic response to a contralateral sound is created by interaction of a sustained excitatory input with a sustained inhibitory input, also from the contralateral ear. Whether the response is to the onset or offset of a sound is determined by the relative timing between the excitatory and inhibitory inputs. Thus, in MSO of the mustached bat, the ipsilateral excitatory pathway from the cochlear nucleus seen in animals with low-frequency hearing is virtually absent, and the MSO is adapted for timing analysis by using input from only the contralateral ear.

Action Potentials↗
PubMed1992

Frequency tuning properties of neurons in the inferior colliculus of an FM bat.

We examined frequency tuning characteristics of single neurons in the inferior colliculus of the echolocating bat, Eptesicus fuscus, in order to determine whether there are different classes of spectral selectivity at this level and to relate frequency tuning properties to the design of the echolocation signal. In unanesthetized but tranquilized animals, we recorded responses from 363 single units to pure tones, frequency-modulated (FM) sweeps, or broad-band noise. Most units were selective for stimulus type; 50% responded only to pure tones, 14% responded only to FM sweeps, and 5% responded only to noise. The remainder responded to two or more types of stimuli. Tuning curves could be classified as follows: 1) V-shaped tuning curves (57%) were the most common type; 2) closed tuning curves (20%) had thresholds at both low and high sound levels; 3) narrow filters (14%) had Q values above 20 at 10 dB and 30 dB above threshold or 10 dB and 40 dB above threshold; 4) frequency-opponent tuning (6%) was found in units with high spontaneous activity; within a center range of frequencies, firing rate increased above spontaneous level, but at higher or lower frequencies, firing rate decreased below spontaneous level; 5) double-tuned units (3%) had two best frequencies (BF). The most clear evidence of topographic distribution was seen for filter units, which were only found in the dorsal part of the 20-30 kHz isofrequency contour. Filter units were also the most clearly related to the echolocation signal of the bat. Their BFs were all within the range of the dominant frequency (approximately 20-30 kHz) that Eptesicus uses during the searching phase of echolocation.

Acoustic Stimulation↗
PubMed1991

Binaural properties of single units in the superior olivary complex of the mustached bat.

1. Previous studies of the superior olive of echolocating bats suggest that the lateral superior olive (LSO) retains the same structure and function as in other mammals but that the medial superior olive (MSO) is different in structure and possibly also in function. The present study is an examination of this idea in Pteronotus parnellii, a bat that has a large and well-defined MSO. 2. Using pure tones presented via earphones, we obtained data on frequency tuning for 60 single units and 96 multiunits in LSO and 94 single units and 154 multiunits in MSO. Of these we also obtained binaural response characteristics from 55 single units in LSO and 72 single units in MSO. 3. LSO and MSO each have a complete tonotopic representation, arranged in a sequence similar to that of other mammals studied. However, in both LSO and MSO there is an expanded representation of the frequencies around 60 kHz, the main frequency component of the bat's echolocation call; there is another expanded representation of the range around 90 kHz, the third harmonic of the call. The expansion of these frequency ranges suggests that the functions of LSO and MSO in Pteronotus are related to echolocation behavior. 4. The binaural characteristics of cells in LSO were essentially the same as those seen in other mammals. Most LSO units (93%) were excited by the ipsilateral ear and inhibited by the contralateral ear. The responses of nearly all LSO units were completely suppressed when the sound level at the two ears was equal. 5. The binaural characteristics of cells in MSO were different from those in nonecholocating mammals. Most MSO units (72%) were excited by the contralateral ear but were neither excited nor inhibited by the ipsilateral ear. Of the remaining units, 21% were excited by the contralateral ear and inhibited by the ipsilateral ear, and only 6% were excited by both ears. 6. The temporal discharge patterns of units in MSO differed from the tonic response pattern seen in LSO. Most MSO units had phasic response patterns, with a few spikes at the onset or offset of the stimulus; the response often changed from ON to OFF depending on stimulus frequency. 7. The results support the idea that in evolution LSO has remained unchanged, whereas MSO has undergone adaptation. The function of LSO in Pteronotus seems to be identical to that in other mammals, i.e., analysis of interaural sound level differences to derive azimuthal location. The function of MSO in Pteronotus must be different from that in nonecholocating mammals.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1991

The monaural nuclei of the lateral lemniscus in an echolocating bat: parallel pathways for analyzing temporal features of sound.

In echolocating bats, three cell groups in the lateral lemniscus are conspicuous for their large size and high degree of differentiation. These cell groups are the intermediate nucleus (INLL), columnar nucleus (VNLLc), and multipolar cell area (VNLLm). All receive projections from the contralateral cochlear nucleus. Previous anatomical studies suggest the hypothesis that these nuclei are important for analyzing the temporal structure of sound. To investigate this possibility, we recorded responses of single units in the INLL, VNLLc, and VNLLm of Eptesicus fuscus. The results show that each cytoarchitectural division contains a complete tonotopic representation. Certain response properties are common to all three nuclei. First, virtually all units are monaural. Second, all are broadly tuned to frequency; their average Q10dB value of 9.1 is considerably lower than values measured in the inferior colliculus of Eptesicus. Third, most units have little or no spontaneous activity. Fourth, all have short integration times, responding robustly to stimuli less than 5 msec in duration. The broad tuning, lack of spontaneous activity, and short integration time all make these neurons well suited for the accurate encoding of temporal information. Although there are many similarities, there are also important differences among nuclei. The clearest evidence of specialization is in VNLLc. Neurons here are more broadly tuned than those in INLL or VNLLm, have no spontaneous activity, and always respond with one spike per stimulus. The latency of the spike is precisely locked to the stimulus onset, with variability from trial to trial as low as 0.03 msec. In addition, the latency remains constant over large variations in frequency or intensity. In INLL and VNLLm, response patterns are about equally distributed between tonic, chopping, and phasic; there are no single-spike constant-latency responses of the type seen in VNLLc, although some choppers and pausers do respond with constant first-spike latency. The results indicate that VNLLc is specialized to encode very precisely the onset of sound; the other nuclei may encode ongoing properties of a sound.

Acoustic Stimulation↗
PubMed1989

Central acoustic tract in an echolocating bat: an extralemniscal auditory pathway to the thalamus.

To determine the sources and targets of auditory pathways that bypass the inferior colliculus in the mustache bat, we injected WGA-HRP in the medial geniculate body and related auditory nuclei of the thalamus as well as in the lower brainstem. We used electrophysiological methods to verify that the injection electrode was in an area responsive to sound. The only thalamic injections that produced retrograde transport to cells in auditory nuclei caudal to the inferior colliculus were those that included the suprageniculate nucleus. These injections labeled a group of large multipolar cells lying between the ventral nucleus of the lateral lemniscus and the superior olivary complex. Neurons in this cell group have also been shown to project to the deep layers of the superior colliculus in the mustache bat. The pathway revealed by these studies is almost identical to the "central acoustic tract" in which fibers course medial to the lateral lemniscus and bypass the inferior colliculus to reach the deep superior colliculus and the suprageniculate nucleus.

Animals↗
PubMed1988

Connections of the superior olivary complex in the rufous horseshoe bat Rhinolophus rouxi.

In the rufous horseshoe bat (Rhinolophus rouxi), the superior olivary complex contains four main divisions. In comparison with other species, the most lateral division is clearly homologous to the lateral superior olive (LSO); the most medial division is homologous to the medial nucleus of the trapezoid body (MNTB). Lying between these landmarks, in approximately the position of the medial superior olive (MSO) of other mammals, are two additional divisions that are cytoarchitecturally distinct from one another yet do not greatly resemble the MSO of nonecholocating mammals such as the cat. We refer to these nuclei as the dorsal medial superior olive (DMSO) and the ventral medial superior olive (VMSO). We examined the afferent and efferent connections of all of these cell groups with retrograde and anterograde transport of WGA-HRP from the superior olivary complex. In the same animals we recorded the binaural response properties of single units in the superior olivary complex. Virtually all units recorded in LSO were excitatory to the ipsilateral ear and inhibitory to the contralateral ear (EI); all of the units sampled in the MNTB and most of those sampled in the VMSO responded only to the contralateral ear (OE). In DMSO the binaural properties of units were varied: the number of units that were inhibitory to the ipsilateral ear and excitatory to the contralateral ear (IE) was about equal to the number of units excitatory to both ears (EE); a few units had OE responses; no units had EI responses. Connectional correlates for these binaural response properties are seen in the patterns of retrograde transport from WGA-HRP injections in the divisions of the superior olive. The LSO receives projections from the ipsilateral cochlear nucleus and MNTB; MNTB receives projections from the contralateral cochlear nucleus. The DMSO and VMSO both receive bilateral projections from the cochlear nuclei. The results of retrograde and anterograde transport suggest that VMSO, in addition, receives projections from the ipsilateral MNTB. The LSO, DMSO, and VMSO all project to the ventral two-thirds of the central nucleus of the inferior colliculus, and their targets are approximately coextensive. However, the LSO projects bilaterally to the inferior colliculus, whereas the medial cell groups project mainly ipsilaterally.

Acoustic Stimulation↗
PubMed1987

Subcortical connections of the superior colliculus in the mustache bat, Pteronotus parnellii.

The mustache bat, Pteronotus parnellii, depends on echolocation to navigate and capture prey. This adaptation is reflected in the large size and elaboration of brainstem auditory structures and in the minimal development of visual structures. The superior colliculus, usually associated with orienting the eyes, is nevertheless large and well developed in Pteronotus. This observation raises the question of whether the superior colliculus in the echolocating bat has evolved to play a major role in auditory rather than visual orientation. The connections of the superior colliculus in Pteronotus were studied with the aid of anterograde and retrograde transport of wheat germ agglutinin conjugated to HRP. These results indicate that the superior colliculus of Pteronotus is composed almost entirely of the layers beneath stratum opticum. The retinal projection is restricted to a very thin zone just beneath the pial surface. Prominent afferent pathways originate in motor structures, particularly the substantia nigra and the deep nuclei of the cerebellum. Sensory input from the auditory system originates in three brainstem nuclei: the inferior colliculus, the anterolateral periolivary nucleus, and the dorsal nuclei of the lateral lemniscus. The projections from these auditory structures terminate mainly in the central tier of the deep layer. The most prominent efferent pathways are those to medial motor structures of the contralateral brainstem via the predorsal bundle and to the ipsilateral midbrain and pontine tegmentum via the lateral efferent bundle. Ascending projections to the diencephalon are mainly to the medial dorsal nucleus and zona incerta. Thus, the superior colliculus in Pteronotus possesses well-developed anatomical connections that could mediate reflexes for orienting its ears, head, or body toward objects detected by echolocation.

Afferent Pathways↗
PubMed1986

Connectional basis for frequency representation in the nuclei of the lateral lemniscus of the bat Eptesicus fuscus.

To study the role of the lateral lemniscus as a link in the ascending auditory pathway, injections of neuronal tracers were placed in the anteroventral cochlear nucleus (AVCN) and in the inferior colliculus of the bat Eptesicus fuscus. To correlate the anatomical results with tonotopic organization, the characteristic frequency of cells at each injection site was determined electrophysiologically. Pathways from AVCN diverge to 3 major targets in the lateral lemniscus, the intermediate nucleus and 2 divisions of the ventral nucleus (VNLL). Projections from these 3 nuclei then converge at the inferior colliculus. One cell group is particularly notable for its cytoarchitectural appearance. It is referred to here as the columnar area of VNLL because its cells are organized as a tightly packed matrix of columns and rows. The connections of the columnar area are organized in sheets that are precisely related to the tonotopic organization of both AVCN and the inferior colliculus. Sheets of cells in the dorsal part of the columnar area receive projections from low-frequency parts of AVCN and project to low-frequency parts of the inferior colliculus. These sheets of connections occupy successively more ventral locations as the tonotopic focus of the injection site increases in frequency. The entire range of frequencies audible to the bat is systematically represented along the dorsal-ventral dimension of the columnar area. Because each column is only 20-30 cells in height, frequency representation must be compressed in this dimension. Within the columnar area there is an overrepresentation of frequencies between 25 and 50 kHz, which corresponds roughly to the range of the FM echo-location call in Eptesicus. The connections of the other nuclei of the lateral lemniscus are not as precisely related to the tonotopic organization of the system as are those of the columnar area.

Afferent Pathways↗
PubMed1984

Projections from the superior olivary complex to the cochlear nucleus in the tree shrew.

The origins and targets of projections from the superior olivary complex to the cochlear nuclei were studied in the tree shrew by placing small injections of horseradish peroxidase (HRP) in the cochlear nucleus and small injections of 3H-leucine in the superior olivary complex. The results show that the descending pathways originate in periolivary cell groups surrounding the medial and lateral superior olives and that periolivary nuclei differ from one another in their patterns of projections to the cochlear nucleus. For example, cell groups may project either ipsilaterally or bilaterally. Cells in the lateral nucleus of the trapezoid body project only to the ipsilateral cochlear nucleus. Other periolivary cell groups project bilaterally, although some of these may project more heavily to one side than the other. Some pathways have widespread targets in the cochlear nucleus whereas others have relatively specific targets. Diffuse projections to all divisions of the cochlear nucleus arise from the lateral nucleus of the trapezoid body ipsilaterally and from the medial perioliviary nucleus bilaterally. The targets of other descending pathways are more restricted. The anterolateral, dorsal, and dorsolateral periolivary nuclei project mainly to the anteroventral cochlear nucleus; the ventral nucleus of the trapezoid body and the posterior periolivary nucleus project mainly to the dorsal and posteroventral cochlear nuclei. All of these specific projections are bilateral. These results suggest that projections from the periolivary cell groups to the cochlear nucleus consist of multiple components with different degrees of specificity.

Animals↗
PubMed1980

Neuron and stimulus typologies in the rat gustatory system.

Although gustatory neurons may be categorized in terms of one or a few characteristics (e.g. 'best stimulus'), such typologies are essentialistic and inconsistent with modern taxonomic methods. If olythetic taxonomic criteria are used and the variability among neuronal responses is closely analyzed, neuronal 'types' are found to disappear, at least within the acid-salt range. This applies to both the primary nerve level (chorda tympani nerve) and secondary level (nucleus tractus solitarius) of the taste system in the rat. In the same context, taste stimuli may fall into different groups if several very similar stimuli are used (e.g. sodium and lithium salts). This is not surprising, and may depend on the choice of stimulus arrays rather than a differentiation of a few stimulus types by the taste system. Finally, it should be noted that the arguments regarding neuron and stimulus typologies presented here for the taste system are also valid for other sensory systems, although the conclusions may be different.

Animals↗
PubMedDate not supplied

Neural population coding and auditory temporal pattern analysis.

Over the 2 decades that have elapsed since Robert Erickson first published his pioneering work on across-fiber patterns in the gustatory system, the idea that information is represented by a population code has become almost universally accepted among neuroscientists. Although the concept of a population code is an implicit theoretical assumption underlying most of the work done in neuroscience today, the details of how population codes operate in specific systems remain unclear in many respects. This article reviews electrophysiological studies of the auditory system of echolocating bats that show that information about sound is initially represented across both space and time by relative amounts of activity in populations of excitatory and inhibitory neurons with different discharge patterns, different sensitivity functions, and different latencies. At the next level, each neuron in the auditory midbrain receives convergent input from a specific population of these lower brainstem neurons and acts as a "readout" of activity within this population. As a result, midbrain neurons become selectively tuned to stimulus features, for example, signal duration, to which neurons at lower levels respond indiscriminately. Intracellular recordings from auditory midbrain neurons show some of the mechanisms by which population input is processed. The known projection patterns of the midbrain "readout" neurons indicate that they, in turn, must become part of a new spatio-temporal population code that is transmitted to neurons at the thalamus, where additional forms of selectivity and patterns of output arise.

Animals↗